Followup to: Evolutions Are Stupid
It is a very common misconception that an evolution works for the good of its species. Can you remember hearing someone talk about two rabbits breeding eight rabbits and thereby "contributing to the survival of their species"? A modern evolutionary biologist would never say such a thing; they'd sooner breed with a rabbit.
It's yet another case where you've got to simultaneously consider multiple abstract concepts and keep them distinct. Evolution doesn't operate on particular individuals; individuals keep whatever genes they're born with. Evolution operates on a reproducing population, a species, over time. There's a natural tendency to think that if an Evolution Fairy is operating on the species, she must be optimizing for the species. But what really changes are the gene frequencies, and frequencies don't increase or decrease according to how much the gene helps the species as a whole. As we shall later see, it's quite possible for a species to evolve to extinction.
Why are boys and girls born in roughly equal numbers? (Leaving aside crazy countries that use artificial gender selection technologies.) To see why this is surprising, consider that 1 male can impregnate 2, 10, or 100 females; it wouldn't seem that you need the same number of males as females to ensure the survival of the species. This is even more surprising in the vast majority of animal species where the male contributes very little to raising the children—humans are extraordinary, even among primates, for their level of paternal investment. Balanced gender ratios are found even in species where the male impregnates the female and vanishes into the mist.
Consider two groups on different sides of a mountain; in group A, each mother gives birth to 2 males and 2 females; in group B, each mother gives birth to 3 females and 1 male. Group A and group B will have the same number of children, but group B will have 50% more grandchildren and 125% more great-grandchildren. You might think this would be a significant evolutionary advantage.
But consider: The rarer males become, the more reproductively valuable they become—not to the group, but to the individual parent. Every child has one male and one female parent. Then in every generation, the total genetic contribution from all males equals the total genetic contribution from all females. The fewer males, the greater the individual genetic contribution per male. If all the females around you are doing what's good for the group, what's good for the species, and birthing 1 male per 10 females, you can make a genetic killing by birthing all males, each of whom will have (on average) ten times as many grandchildren as their female cousins.
So while group selection ought to favor more girls, individual selection favors equal investment in male and female offspring. Looking at the statistics of a maternity ward, you can see at a glance that the quantitative balance between group selection forces and individual selection forces is overwhelmingly tilted in favor of individual selection in Homo sapiens.
(Technically, this isn't quite a glance. Individual selection favors equal parental investments in male and female offspring. If males cost half as much to birth and/or raise, twice as many males as females will be born at the evolutionarily stable equilibrium. If the same number of males and females were born in the population at large, but males were twice as cheap to birth, then you could again make a genetic killing by birthing more males. So the maternity ward should reflect the balance of parental opportunity costs, in a hunter-gatherer society, between raising boys and raising girls; and you'd have to assess that somehow. But ya know, it doesn't seem all that much more reproductive-opportunity-costly for a hunter-gatherer family to raise a girl, so it's kinda suspicious that around the same number of boys are born as girls.)
Natural selection isn't about groups, or species, or even individuals. In a sexual species, an individual organism doesn't evolve; it keeps whatever genes it's born with. An individual is a once-off collection of genes that will never reappear; how can you select on that? When you consider that nearly all of your ancestors are dead, it's clear that "survival of the fittest" is a tremendous misnomer. "Replication of the fitter" would be more accurate, although technically, fitness is defined only in terms of replication.
Natural selection is really about gene frequencies. To get a complex adaptation, a machine with multiple dependent parts, each new gene as it evolves depends on the other genes being reliably present in its genetic environment. They must have high frequencies. The more complex the machine, the higher the frequencies must be. The signature of natural selection occurring is a gene rising from 0.00001% of the gene pool to 99% of the gene pool. This is the information, in an information-theoretic sense; and this is what must happen for large complex adaptations to evolve.
The real struggle in natural selection is not the competition of organisms for resources; this is an ephemeral thing when all the participants will vanish in another generation. The real struggle is the competition of alleles for frequency in the gene pool. This is the lasting consequence that creates lasting information. The two rams bellowing and locking horns are only passing shadows.
It's perfectly possible for an allele to spread to fixation by outcompeting an alternative allele which was "better for the species". If the Flying Spaghetti Monster magically created a species whose gender mix was perfectly optimized to ensure the survival of the species—the optimal gender mix to bounce back reliably from near-extinction events, adapt to new niches, etcetera—then the evolution would rapidly degrade this species optimum back into the individual-selection optimum of equal parental investment in males and females.
Imagine a "Frodo gene" that sacrifices its vehicle to save its entire species from an extinction event. What happens to the allele frequency as a result? It goes down. Kthxbye.
If species-level extinction threats occur regularly (call this a "Buffy environment") then the Frodo gene will systematically decrease in frequency and vanish, and soon thereafter, so will the species. A hypothetical example? Maybe. If the human species was going to stay biological for another century, it would be a good idea to start cloning Gandhi.
In viruses, there's the tension between individual viruses replicating as fast as possible, versus the benefit of leaving the host alive long enough to transmit the illness. This is a good real-world example of group selection, and if the virus evolves to a point on the fitness landscape where the group selection pressures fail to overcome individual pressures, the virus could vanish shortly thereafter. I don't know if a disease has ever been caught in the act of evolving to extinction, but it's probably happened any number of times.
Segregation-distorters subvert the mechanisms that usually guarantee fairness of sexual reproduction. For example, there is a segregation-distorter on the male sex chromosome of some mice which causes only male children to be born, all carrying the segregation-distorter. Then these males impregnate females, who give birth to only male children, and so on. You might cry "This is cheating!" but that's a human perspective; the reproductive fitness of this allele is extremely high, since it produces twice as many copies of itself in the succeeding generation as its nonmutant alternative. Even as females become rarer and rarer, males carrying this gene are no less likely to mate than any other male, and so the segregation-distorter remains twice as fit as its alternative allele. It's speculated that real-world group selection may have played a role in keeping the frequency of this gene as low as it seems to be. In which case, if mice were to evolve the ability to fly and migrate for the winter, they would probably form a single reproductive population, and would evolve to extinction as the segregation-distorter evolved to fixation.
Around 50% of the total genome of maize consists of transposons, DNA elements whose primary function is to copy themselves into other locations of DNA. A class of transposons called "P elements" seem to have first appeared in Drosophila only in the middle of the 20th century, and spread to every population of the species within 50 years. The "Alu sequence" in humans, a 300-base transposon, is repeated between 300,000 and a million times in the human genome. This may not extinguish a species, but it doesn't help it; transposons cause more mutations which are as always mostly harmful, decrease the effective copying fidelity of DNA. Yet such cheaters are extremely fit.
Suppose that in some sexually reproducing species, a perfect DNA-copying mechanism is invented. Since most mutations are detrimental, this gene complex is an advantage to its holders. Now you might wonder about beneficial mutations—they do happen occasionally, so wouldn't the unmutable be at a disadvantage? But in a sexual species, a beneficial mutation that began in a mutable can spread to the descendants of unmutables as well. The mutables suffer from degenerate mutations in each generation; and the unmutables can sexually acquire, and thereby benefit from, any beneficial mutations that occur in the mutables. Thus the mutables have a pure disadvantage. The perfect DNA-copying mechanism rises in frequency to fixation. Ten thousand years later there's an ice age and the species goes out of business. It evolved to extinction.
The "bystander effect" is that, when someone is in trouble, solitary individuals are more likely to intervene than groups. A college student apparently having an epileptic seizure was helped 85% of the time by a single bystander, and 31% of the time by five bystanders. I speculate that even if the kinship relation in a hunter-gatherer tribe was strong enough to create a selection pressure for helping individuals not directly related, when several potential helpers were present, a genetic arms race might occur to be the last one to step forward. Everyone delays, hoping that someone else will do it. Humanity is facing multiple species-level extinction threats right now, and I gotta tell ya, there ain't a lot of people steppin' forward. If we lose this fight because virtually no one showed up on the battlefield, then—like a probably-large number of species which we don't see around today—we will have evolved to extinction.
Cancerous cells do pretty well in the body, prospering and amassing more resources, far outcompeting their more obedient counterparts. For a while.
Multicellular organisms can only exist because they've evolved powerful internal mechanisms to outlaw evolution. If the cells start evolving, they rapidly evolve to extinction: the organism dies.
So praise not evolution for the solicitous concern it shows for the individual; nearly all of your ancestors are dead. Praise not evolution for the solicitous concern it shows for a species; no one has ever found a complex adaptation which can only be interpreted as operating to preserve a species, and the mathematics would seem to indicate that this is virtually impossible. Indeed, it's perfectly possible for a species to evolve to extinction. Humanity may be finishing up the process right now. You can't even praise evolution for the solicitous concern it shows for genes; the battle between two alternative alleles at the same location is a zero-sum game for frequency.
Fitness is not always your friend.
Part of the sequence The Simple Math of Evolution
Next post: "The Tragedy of Group Selectionism"
Previous post: "No Evolutions for Corporations or Nanodevices"